C. Dart: Institute of Integrative Biology, University of Liverpool, Crown Street, Liverpool L69 7ZB, UK.
J Physiol. 2013 Oct 15;591(20):5107-23. doi: 10.1113/jphysiol.2013.262006. Epub 2013 Aug 19.
Vasodilator-induced elevation of intracellular cyclic AMP (cAMP) is a central mechanism governing arterial relaxation but is incompletely understood due to the diversity of cAMP effectors. Here we investigate the role of the novel cAMP effector exchange protein directly activated by cAMP (Epac) in mediating vasorelaxation in rat mesenteric arteries. In myography experiments, the Epac-selective cAMP analogue 8-pCPT-2-O-Me-cAMP-AM (5 μM, subsequently referred to as 8-pCPT-AM) elicited a 77.6 ± 7.1% relaxation of phenylephrine-contracted arteries over a 5 min period (mean ± SEM; n = 6). 8-pCPT-AM induced only a 16.7 ± 2.4% relaxation in arteries pre-contracted with high extracellular K(+) over the same time period (n = 10), suggesting that some of Epac's relaxant effect relies upon vascular cell hyperpolarization. This involves Ca(2+)-sensitive, large-conductance K(+) (BK(Ca)) channel opening as iberiotoxin (100 nM) significantly reduced the ability of 8-pCPT-AM to reverse phenylephrine-induced contraction (arteries relaxed by only 35.0 ± 8.5% over a 5 min exposure to 8-pCPT-AM, n = 5; P < 0.05). 8-pCPT-AM increased Ca(2+) spark frequency in Fluo-4-AM-loaded mesenteric myocytes from 0.045 ± 0.008 to 0.103 ± 0.022 sparks s(-1) μm(-1) (P < 0.05) and reversibly increased both the frequency (0.94 ± 0.25 to 2.30 ± 0.72 s(-1)) and amplitude (23.9 ± 3.3 to 35.8 ± 7.7 pA) of spontaneous transient outward currents (STOCs) recorded in isolated mesenteric myocytes (n = 7; P < 0.05). 8-pCPT-AM-activated STOCs were sensitive to iberiotoxin (100 nM) and to ryanodine (30 μM). Current clamp recordings of isolated myocytes showed a 7.9 ± 1.0 mV (n = 10) hyperpolarization in response to 8-pCPT-AM that was sensitive to iberiotoxin (n = 5). Endothelial disruption suppressed 8-pCPT-AM-mediated relaxation in phenylephrine-contracted arteries (24.8 ± 4.9% relaxation after 5 min of exposure, n = 5; P < 0.05), as did apamin and TRAM-34, blockers of Ca(2+)-sensitive, small- and intermediate-conductance K(+) (SK(Ca) and IK(Ca)) channels, respectively, and N(G)-nitro-L-arginine methyl ester, an inhibitor of nitric oxide synthase (NOS). In Fluo-4-AM-loaded mesenteric endothelial cells, 8-pCPT-AM induced a sustained increase in global Ca(2+). Our data suggest that Epac hyperpolarizes smooth muscle by (1) increasing localized Ca(2+) release from ryanodine receptors (Ca(2+) sparks) to activate BK(Ca) channels, and (2) endothelial-dependent mechanisms involving the activation of SK(Ca)/IK(Ca) channels and NOS. Epac-mediated smooth muscle hyperpolarization will limit Ca(2+) entry via voltage-sensitive Ca(2+) channels and represents a novel mechanism of arterial relaxation.
血管扩张剂诱导细胞内环腺苷酸 (cAMP) 的升高是调节动脉舒张的核心机制,但由于 cAMP 效应物的多样性,其机制尚不完全清楚。在这里,我们研究了新型 cAMP 效应物——cAMP 直接激活的交换蛋白 (Epac) 在介导大鼠肠系膜动脉舒张中的作用。在肌动描记法实验中,Epac 选择性 cAMP 类似物 8-pCPT-2-O-Me-cAMP-AM(5 μM,以下简称 8-pCPT-AM)在 5 分钟内引起去氧肾上腺素收缩的血管舒张 77.6±7.1%(平均值±SEM;n=6)。8-pCPT-AM 在相同时间内对高细胞外 K+预收缩的血管仅引起 16.7±2.4%的舒张(n=10),这表明 Epac 的部分舒张作用依赖于血管细胞的超极化。这涉及钙敏感、大电导钾(BK(Ca))通道的开放,因为蜂毒素(100 nM)显著降低了 8-pCPT-AM 逆转去氧肾上腺素诱导收缩的能力(8-pCPT-AM 暴露 5 分钟后,血管仅舒张 35.0±8.5%,n=5;P<0.05)。8-pCPT-AM 增加了 Fluo-4-AM 加载的肠系膜肌细胞中的 Ca2+火花频率,从 0.045±0.008 增加到 0.103±0.022 火花 s-1μm-1(P<0.05),并可逆地增加了分离的肠系膜肌细胞中自发性瞬时外向电流(STOC)的频率(0.94±0.25 增加到 2.30±0.72 s-1)和幅度(23.9±3.3 增加到 35.8±7.7 pA)(n=7;P<0.05)。8-pCPT-AM 激活的 STOC 对蜂毒素(100 nM)和 Ryanodine(30 μM)敏感。分离肌细胞的电流钳记录显示,8-pCPT-AM 引起 7.9±1.0 mV(n=10)的超极化,对蜂毒素敏感(n=5)。内皮破坏抑制了去氧肾上腺素收缩的血管 8-pCPT-AM 介导的舒张(5 分钟暴露后 24.8±4.9%的舒张,n=5;P<0.05),蜂毒素、apamin 和 TRAM-34 分别为钙敏感、小和中电导钾(SK(Ca)和 IK(Ca))通道的阻断剂,N(G)-硝基-L-精氨酸甲酯,一氧化氮合酶(NOS)的抑制剂。在 Fluo-4-AM 加载的肠系膜内皮细胞中,8-pCPT-AM 诱导了持续的全局 Ca2+增加。我们的数据表明,Epac 通过以下两种机制使平滑肌超极化:(1)通过增加肌浆网ryanodine 受体(Ca2+火花)的局部 Ca2+释放来激活 BK(Ca)通道;(2)内皮依赖性机制涉及 SK(Ca)/IK(Ca)通道和 NOS 的激活。Epac 介导的平滑肌超极化将通过电压敏感性 Ca2+通道限制 Ca2+内流,代表动脉舒张的一种新机制。